Chlorophyll Maximum Research Articles

Wind‐driven currents and water masses shape spring phytoplankton distribution and composition in hydrologically complex, productive shelf waters

AbstractThe Northern Patagonian shelf (35–42°S SW South Atlantic) is a transitional area between the subtropical and subantarctic domains, where the shelf shifts from source to sink of atmospheric CO2. In the southeast portion (39–42°S, 61–55°W) of this area, winds interact with different water masses and picture a complex circulation that supports two productive frontal systems: the Mid‐Shelf Front (MSF) and the Patagonian Shelf‐break Front. We combined remote sensing (e.g., monthly surface chlorophyll a over the period: September–December 2002–2021), reanalysis (winds and surface currents), and in situ measurements from three cross‐shelf cruises in spring 2016, 2017, and 2019, to assess the underlying drivers of phytoplankton distribution. The anomalous high chlorophyll concentration in the MSF in spring 2016 was related to enhanced water mass retention by N‐NE winds. Contrasting phytoplankton assemblages emerged related to stratified waters in the shelf influenced by the Patagonian Current and vertically mixed waters at the continental edge (Malvinas Current). Large diatoms prevailed in the outer shelf followed by dinoflagellates, while nanoflagellates and microzooplankton dominated in the mid‐shelf. Furthermore, thickness of the subsurface chlorophyll maximum was broader in the early spring than in a more advanced state of the bloom where the coccolithophore Emiliania huxleyi was abundant. We showed that different time‐scale physical processes—winds, surface currents and vertical mixing—are underpinning the development of spring blooms with different taxa composition in the Patagonian shelf, where different water masses define the hydrological complexity that led to patchy phytoplankton.

Paclobutrazol and maleic hydrazide-induced growth inhibition in warm season turfgrasses through structural and physiological differences

Warm season turfgrasses are fast growing and require frequent mowing, which increases maintenance cost. Various plant growth retardants are used to reduce the growth and required amount of mowing of turfgrasses. Therefore, the present study was planned to explore the impact of paclobutrazol (PBZ) and maleic hydrazide (MH) on the growth rate and quality of warm season turfgrasses. Four turfgrasses were grown in pots at research area, MNS-University of Agriculture, Multan. Paclobutrazol (PBZ) and Maleic Hydrazide at 0.01%, 0.02%, 0.04% levels were applied through foliar application with three repeats in CRD arrangement. The data of growth, quality and anatomical attributes were evaluated. The results showed significant differences in growth and the quality of turfgrasses in response to different levels of plant growth retardants. PBZ application at 0.04% maximum reduced fresh (26%, 35%, 25%) and dry (45%, 54%, 34%) weights of shoot, root and clippings respectively in Fine Dacca turfgrass. Additionally, PBZ application decreased the amount of mowing (40%) compared to control. Fine Dacca also produced the highest visual quality turf with maximum Chlorophyll a (46.87%) and Chlorophyll b (60%) in comparison to nontreated turfgrasses. The minimum number and density of stomatal and mesophyll cells by 45%, 47% were also recorded in Fine Dacca at PBZ (0.04%) with respect to control. It is concluded that PBZ at 0.04% could be used to reduce the amount of mowing in warm season turfgrasses, which will decrease the cost of lawn maintenance and support the landscape industry.

Top-down and bottom-up control of phytoplankton in a mid-latitude continental shelf ecosystem

It is widely recognized that ecosystems are influenced by both bottom-up (resource-driven) and top-down (predation-driven) controls. However, in the context of marine phytoplankton, there is often a strong emphasis on resource availability, with less attention given to the significant impact of top-down control on phytoplankton patterns. Studies in the Gulf of Cádiz have mainly focused on resource control perspectives, overlooking the role of zooplankton predation as a driving force in phytoplankton dynamics. This limitation in research approaches has resulted in an incomplete understanding of phytoplankton biomass patterns, as they cannot be fully explained by resource control alone. This study makes a first attempt at studying the combined bottom-up and top-down controls of phytoplankton in the Gulf of Cádiz and aims to test their relative influence on the variance of phytoplankton biomass. For this purpose, monthly samples of chlorophyll-a, mesozooplankton and environmental variables determining phytoplankton growth were taken over a six-year period. The variables involved in phytoplankton growth were compiled into a single factor (primary production limiting factor, pplf) representing the bottom-up control. Grazing (G) by mesozooplankton represented the top-down control of phytoplankton. Different statistical tools (multiple linear regression, decision trees and random forest) were used to analyse the relative importance of top-down vs. bottom-up control on the variance of phytoplankton biomass (using chlorophyll-a as a proxy). All three techniques found that the top-down control had more relative weight on the variance of phytoplankton biomass for the continental shelf as a whole. Resource availability (i.e., bottom-up control) determined phytoplankton growth rate, although grazing (i.e., top-down control) defined the maximum chlorophyll values in the studied region. Additionally, the results revealed a spatiotemporal pattern in the relative importance of top-down and bottom-up control on phytoplankton dynamics. These findings underscore the significance of considering the impact of top-down control for a comprehensive understanding of phytoplankton dynamics in the Gulf of Cádiz.

Spatiotemporal variations of the oxycline and its response to subduction events in the Arabian Sea

The Arabian Sea is a significant hypoxic region in world’s oceans, characterized by the most extensive oxygen minimum zones (OMZs). Both physical and biological processes can alter the vertical and horizontal distribution of dissolved oxygen within the upper ocean and affect the spatial and temporal distribution of hypoxia within the OMZ. To identify the key physical and biological factors influencing the boundaries of oxycline, we analyzed an extensive dataset collected from the biogeochemical-Argo (BGC-Argo) floats during the period of 2010–2022. In particular, we investigated the impact of physical subduction events on the oxycline. Our results shows that the upper boundary of the oxycline deepened in summer and winter, and seemed to be controlled by the mixed layer depth. In contrast, it was shallower during spring and autumn, mainly regulated by the deep chlorophyll maximum. The lower boundary of the oxycline in the western Arabian Sea was predominantly controlled by regional upwelling and downwelling, as well as Rossby waves in the eastern Arabian Sea. Subduction patches originated from the Arabian Sea High Salinity Water (ASHSW) were observed from the BGC-Argo data, which were found to deepen the lower boundary of the oxycline, and increase the oxygen inventory within the oxycline by 8.3%, leading to a partial decrease in hypoxia levels.

Metagenomic Analyses Reveal the Influence of Depth Layers on Marine Biodiversity on Tropical and Subtropical Regions.

The emergence of open ocean global-scale studies provided important information about the genomics of oceanic microbial communities. Metagenomic analyses shed light on the structure of marine habitats, unraveling the biodiversity of different water masses. Many biological and environmental factors can contribute to marine organism composition, such as depth. However, much remains unknown about microbial communities' taxonomic and functional features in different water layer depths. Here, we performed a metagenomic analysis of 76 publicly available samples from the Tara Ocean Project, distributed in 8 collection stations located in tropical or subtropical regions, and sampled from three layers of depth (surface water layer-SRF, deep chlorophyll maximum layer-DCM, and mesopelagic zone-MES). The SRF and DCM depth layers are similar in abundance and diversity, while the MES layer presents greater diversity than the other layers. Diversity clustering analysis shows differences regarding the taxonomic content of samples. At the domain level, bacteria prevail in most samples, and the MES layer presents the highest proportion of archaea among all samples. Taken together, our results indicate that the depth layer influences microbial sample composition and diversity.

Microbial dynamics in shallow CO2 seeps system off Panarea Island (Italy)

Shallow-water hydrothermal vents are extreme environments characterized by high temperatures, low pH, and high CO2 concentrations; therefore, they are considered as suitable laboratories for studying the effect of global changes on marine microbes. We hypothesized a direct effect of vents on prokaryotic community structure and functioning in the Panarea Island’s hydrothermal system. Sampling was conducted along a 9-station transect characterized by three active emission points. The water column was stratified with a thermocline at 25 m depth and a deep chlorophyll maximum between 50 and 100 m. Prokaryotic abundance ranged from 0.2 to 1.5 × 109 cells L−1, prokaryotic carbon production from 2.4 to 75.4 ng C L−1 h−1, and exoenzymatic activities degrading proteins, phosphorylated compounds, and polysaccharides were on the order of 4–28, 2–31 and 0.2–4.16 nM h−1, respectively. While microbial abundance and production were shaped by the water column's physical structure, alkaline phosphatase and beta-glucosidase activities seemed to be enhanced by hydrothermal fluids. The 16S rRNA gene amplicon sequencing analysis identified a surface, a deep, and a vent-influenced microbial community. In terms of relative abundance members of the SAR11 group dominated the water column, alongside Synechococcus and Prochlorococcus in surface and bottom samples, respectively. Vent-influenced stations were characterized by the presence of Thiomicrorhabdus, a sulfur-oxidizer chemolithoautotroph. Overall, this study provides insights on the coupling between microbial community structure and the biogeochemical cycling of nutrients in low-pH conditions (CO2 and H2S-based), thus addressing some of the opened questions about the response of microbes to acidification.

Zooplankton biomass, size structure, and associated metabolic fluxes with focus on its roles at the chlorophyll maximum layer during the plankton-contaminant MERITE-HIPPOCAMPE cruise

Recent studies have demonstrated that plankton can be a key pathway for the uptake and transfer of contaminants entering the marine environment up to top predators.The plankton-contaminant MERITE-HIPPOCAMPE cruise was devoted to quantifying contaminants in water and the whole plankton size range (10 size fractions) at 10 stations along a north-south transect in the western Mediterranean Sea from the French to the Tunisian coasts through the Provençal and Algerian basins. Pumping and filtering devices and net sampling have been used for collecting very high amounts of small particles and planktonic organisms in the chlorophyll maximum layer (CML). The present paper characterizes the zooplankton components for which the contaminant measurements were carried out. At each station, a horizontal towed Hydro-Bios net with a 60 μm mesh-size net was used to discriminate 5 size-fractions from 60 μm to a few mm. For each size-fraction, one part of the sample was used for dry weight measurements and the other one for estimating the contribution to biomass of detritus, phytoplankton, and among zooplankton of the major taxonomic groups based on the imagery tools ZOOSCAN and FLOWCAM. In each zooplankton size fraction, metabolic rates were calculated from the size spectrum to estimate trophic and excretion fluxes flowing through this fraction. These observations were compared to a similar analysis of tows in the upper layer (vertical) and the surface layer (horizontal).The total sampled biomass concentration at the CML was higher than in the water column (COL) and much higher than at the surface (SURF) in most of the stations, but in the CML and COL a substantial contribution was due to detritus mostly concentrated in the smallest size-fractions (60–200 μm and 200–500 μm). Absolute values of zooplankton biomass show neither a clear spatial pattern nor a significant difference between strata. The CML layer was dominated by copepods similarly to COL and SURF, but presented a higher contribution of nauplii and a near absence of appendicularians. At some stations, crustaceans and gelatinous plankton could be important contributors to CML. The zooplankton biomass composition of the two smallest fractions (<500 μm) was dominated by nauplii, small copepods and, occasionally, by small miscellaneous organisms (mostly pteropodes). In contrast, clear differences between stations appeared for the largest fractions (>500 μm) due to large crustaceans, gelatinous organisms, and chaetognaths. These changes in biomass composition according to size fractions suggest a progressive trophic shift from dominant herbivory in the smallest fractions to more contrasted trophic structure (including carnivory) in the largest fractions.The daily carbon demand and the N and P excretion of zooplankton were on average higher at the CML but with no significant difference with COL. The zooplankton grazing represented 2.7 to 22.7 % of the phytoplankton stock per day, whereas its excretion represented a daily N and P recycling compared to dissolved inorganic nitrogen and phosphorus stocks ranging respectively from 0.2 to 19 % and from 0 to 21 %. This information should help in the interpretation of the content of various contaminants in zooplankton fractions.

Phytoplankton Diversity and Co-Dependency in a Stratified Oligotrophic Ecosystem in the South Adriatic Sea

The oligotrophy of the southern Adriatic Sea is characterized by seasonal stratification which enables nutrient supply to the euphotic layer. A set of interdisciplinary methods was used to elucidate the diversity and co-dependency of bacterio- and phytoplankton of the water column during the stratification period of July 2021. A total of 95 taxa were determined by microscopy: 58 diatoms, 27 dinoflagellates, 6 coccolithophores, and 4 other autotrophs, which included Chlorophyceae, Chrysophyceae, and Cryptophytes. Nanophytoplankton abundances were higher in comparison to microphytoplankton. The prokaryotic plankton community as revealed by HTS was dominated by Proteobacteria (41–73%), Bacteroidota (9.5–27%), and cyanobacteria (1–10%), while the eukaryotic plankton community was composed of parasitic Syndiniales (45–80%), Ochrophyta (2–18%), Ciliophora (2–21%), Chlorophytes (2–4%), Haptophytes (1–4%), Bacillariophyta (1–13%), Pelagophyta (0.5–12%) and Chrysophyta (0.5–3%). Flow cytometry analysis has recorded Prochlorococcus and photosynthetic picoeukaryotes as more abundant in deep chlorophyll maximum (DCM), and Synechococcus and heterotrophic bacteria as most abundant in surface and thermocline layers. Surface, thermocline, and DCM layers were distinct considering community diversity, temperature, and nutrient correlations, while extreme nutrient values at the beginning of the investigating period indicated a possible nutrient flux. Nutrient and temperature were recognized as the main environmental drivers of phytoplankton and bacterioplankton community abundance.

Phycotoxins in bivalves from the western Canadian Arctic: The first evidence of toxigenicity

This study presents the first evidence that a diverse suite of phycotoxins is not only being actively produced by the toxigenic algal communities in the Canadian Arctic waters, but is also entering the marine food web. We detected measurable amounts of Amnesic Shellfish Toxins (ASTs) and Paralytic Shellfish Toxins (PSTs), as well as trace amounts of other lipophilic toxin groups including pectenotoxins, yessotoxins, and cyclic imines, in bivalves collected from the Canadian Beaufort Sea in 2014 and 2018. There appear to be species-specific differences in accumulation and retention of AST by Arctic bivalves, with significantly higher concentrations recorded in Nuculanidae than Propeamussiidae, likely reflecting physiological and allometric differences. We further confirm the omnipresence of potentially toxic taxonomically-versatile phytoplankton communities in the western Canadian Arctic comprising Pseudo-nitzschia delicatissima group, P. obtusa, Dinophysis acuminata, Prorocentrum minimum, Alexandrium tamarense, and Gymnodinium spp. Although measurements of actual toxicity levels and profiles of these species at the time of sampling fall outside of the scope of this study, we show that high abundance and competitive success of known AST-producers, Pseudo-nitzschia spp., are possible in Canadian Arctic waters. In 2014, a strong dominance of Pseudo-nitzschia spp. was observed at a few shallow coastal stations, representing nearly 40% of the total phytoplankton cell abundances with > 106 cells/L at the depth of maximum chlorophyll a. We further describe oceanographic conditions conducive to high abundances of toxin-producing algae, indicating that temperature is likely a key factor. Even though measured AST and PST concentrations in bivalve tissue remained well below the Health Canada's levels at which monitored fisheries would close, i.e., 5% and 4%, respectively, their presence demonstrate that phycotoxin accumulation is occurring in food webs of the Canadian Beaufort Sea. Yet, the phycotoxin production controls and trophic transfer mechanisms remain unknown. Canadian Arctic marine ecosystems are rapidly changing and temperatures are expected to continue to increase. Given that these changes simultaneously affect multiple, and often co-occurring, species of primary producers, adaptive capacity is likely to play an important role in the structure of phytoplankton communities in the Canadian Arctic.

The Bay of Bengal exposes abundant photosynthetic picoplankton and newfound diversity along salinity-driven gradients.

The Bay of Bengal (BoB) is a 2,600,000 km2 expanse in the Indian Ocean upon which many humans rely. However, the primary producers underpinning food chains here remain poorly characterized. We examined phytoplankton abundance and diversity along strong BoB latitudinal and vertical salinity gradients-which have low temperature variation (27-29°C) between the surface and subsurface chlorophyll maximum (SCM). In surface waters, Prochlorococcus averaged 11.7 ± 4.4 × 104 cells ml-1 , predominantly HLII, whereas LLII and 'rare' ecotypes, HLVI and LLVII, dominated in the SCM. Synechococcus averaged 8.4 ± 2.3 × 104 cells ml-1 in the surface, declined rapidly with depth, and population structure of dominant Clade II differed between surface and SCM; Clade X was notable at both depths. Across all sites, Ostreococcus Clade OII dominated SCM eukaryotes whereas communities differentiated strongly moving from Arabian Sea-influenced high salinity (southerly; prasinophytes) to freshwater-influenced low salinity (northerly; stramenopiles, specifically, diatoms, pelagophytes, and dictyochophytes, plus the prasinophyte Micromonas) surface waters. Eukaryotic phytoplankton peaked in the south (1.9 × 104 cells ml-1 , surface) where a novel Ostreococcus was revealed, named here Ostreococcus bengalensis. We expose dominance of a single picoeukaryote and hitherto 'rare' picocyanobacteria at depth in this complex ecosystem where studies suggest picoplankton are replacing larger phytoplankton due to climate change.

Kedalaman Klorofil Maksimum Selama Musim Peralihan II dan Implikasinya untuk Perikanan Tuna di Laut Banda

The Deep chlorophyll maximum (DCM) in the Banda Sea during the second transitional season (October 2015) was relatively shallow (25 - 64 m) above the mixed layer and far from the base of the euphotic layer or compensation depth. Its presence was identical to the depth of nitraclin injected into the mixed layer as a result of upwelling generated by internal waves. The contribution of these waves to the increase in Banda Sea fertility calculated using ocean color data is 56.87%. This phenomenon is due to the shallow optical depth of the satellite (20-25 m) while high chlorophyll-a concentrations are below. DCM was also associated with the swimming layer of Madidihang (Thunnus albacares) (34 - 68 m) caught with longline gear during the same period.

Ameliorative effects of microbial fertiliser on yield and quality parameters of curly lettuce and cucumber with fertiliser saving

ABSTRACT This research aimed to evaluate the potential effects of microbial fertilizer on the growth, yield and quality parameters and fertiliser savings of lettuce and cucumber. The study had control (C), microbial fertilisation (MF), chemical fertilisation (CF), chemical fertilisation + microbial fertilisation (CF + MF), 50% chemical fertilisation + microbial fertilisation (50% CF + MF) and dipping into microbial fertiliser + chemical fertilisation (D + CF) treatments. The highest head length, root collar diameter, soluble solids and leaf number of lettuce were obtained in D+CF. Maximum chlorophyll contents were obtained in CF, CF + MF and D + CF, all of which were higher than the C application. D + CF and CF + MF resulted in the highest total and marketable yields. In cucumber cultivation, although the highest fruit length was obtained in CF, CF + MF, 50% CF + MF and D + CF were only slightly less than CF. The highest fruit diameters were in CF + MF and 50% CF + MF. The chlorophyll contents were significantly more in 50% CF + MF and D + CF than in other applications. The highest first- and second-class yields were in 50% CF + MF and were significantly higher than those in C and CF. Average fruit weight was also higher in all MF applications. It counclud that the alone MF application is not important, so MF has to be used with organic or chemical fertilisers in lettuce and cucumber cultivation.

Distinct vertical behavior of key Arctic copepods following the midnight sun period in the East Siberian continental margin region, Arctic Ocean

Diel vertical migration (DVM) of zooplankton plays a vital role in biological carbon pump and food web interactions. However, there is considerable debate about the DVM of zooplankton in response to environmental changes in the Arctic Ocean. We investigated DVM behavior in the key Arctic copepods Calanus glacialis, Calanus hyperboreus, and Metridia longa following the midnight sun period in the East Siberian continental margin region. The two Calanus species showed non-DVM behaviors, whereas M. longa showed a typical DVM pattern consistent with the solar radiation cycle. Additionally, these species showed different vertical distributions. Calanus glacialis was distributed at depths above 20 m in the warm fresh water, where the highest density gradient was observed. Calanus hyperboreus was distributed at depths between 30 and 55 m in the cold salty water, where a high contribution of micro phytoplankton and the subsurface chlorophyll maximum (SCM) layer were observed. M. longa was found across a broader range of temperature and salinity than both Calanus species, and it was distributed in the upper water column, where the SCM layer was observed at night and at depths between 100 and 135 m in the daytime. These results imply that M. longa can be well adapted to the changing Arctic Ocean environment, where sea ice loss and ocean warming are ongoing, whereas C. hyperboreus can be the most vulnerable to these changes. These findings provide important information for understanding variations in the vertical distributions of key copepod species in the rapidly changing Arctic marine environment.

Hydrography and deep chlorophyll maximum patterns of the Athos Basin and the Thracian Sea continental shelf (North Aegean Sea) combining glider and satellite observations

This work focuses on the North Aegean Sea deep chlorophyll maximum (DCM) dynamics, as revealed during the first glider mission over the continental shelf and slope of the Thracian Sea and along the Athos basin, and presents a conceptual model on the response of the DCM in relation to the prevailing stratification/mixing of the water column. Water temperature, conductivity, salinity, chlorophyll-a, CDOM, DO and particle backscatter data (bb700), were recorded during this underwater glider mission in summer 2019. Daily satellite data from Sentinel 3 complemented the glider collected data to validate the satellite-derived estimates of surface Chlorophyll-a levels. The high-resolution physical and chemical observations were analyzed emphasizing on thermal stratification, pycnocline strength, CDOM and dissolved oxygen vertical distribution and the presence of deep chlorophyll maximum (DCM). DCM statistical characteristics were derived by fitting a Gaussian model on the chlorophyll-a profiles. The mean DCM was observed at 76 ± 14.5 m depth, located below the euphotic layer, estimated at 64 ± 2.1 m, with the water column dynamics affecting directly its depth, thickness and peak prominence. The satellite-derived chlorophyll-a, integrated within the euphotic layer, was found in fair agreement to the integrated chlorophyll-a values from surface to optical depth recorded by the glider. Chl-a at the DCM were found up to 0.60 mg m−3, the vertically-integrated pigment content was estimated at 45 ± 14 mg m−2 and the values of Optical Community Index (OCI) illustrate the dominance of small-sized plankton. The increased variability in the particulate backscattering coefficient bb700 in Thracian Sea shelf and slope deep waters seems related to sediment resuspension. CDOM levels increased from 0.76 ppb at the sea surface to 1.11 ppb at 40 m depth, being positively correlated to Chl-a within the DCM. CDOM concentration increased steadily at intermediate and deeper waters up to 1.34 ± 0.07 ppb. DO profiles revealed the presence of an oxycline in the DCM layer, capable to oxygenate the intermediate and deeper waters (+3 × 10−3 ml/L/m). Bottom DO in Athos basin was measured at 5.0 ml L−1, indicating that the apparent oxygen utilization in Athos Basin is approximately 50 μmol L−1.

Hypoxia formation triggered by the organic matter from subsurface chlorophyll maximum in a large estuary-shelf system

This study reports, for the first time, the role of shoreward transport of organic matter (OM) from subsurface chlorophyll maximum (SCM) in triggering hypoxia off the Pearl River Estuary (PRE, an outstanding example of typical estuary-shelf systems) based on field measurements. Compared to frequently observed hypoxia driven by surface eutrophication and terrestrial OM during large river discharge, we demonstrate that the upslope-transported SCM played a critical role in forming offshore hypoxia during low river discharge. Together with the plume-sourced OM trapped below the surface plume front, upslope-transported OM originating from the SCM accumulated underneath the pycnocline and consumed dissolved oxygen (DO), enhancing the bottom hypoxia. The DO consumption induced by the SCM-associated OM was estimated to contribute ∼ 26% (±23%) of the DO depletion under the pycnocline. Based on coherent and consistent physical and biogeochemical evidence and reasoning, this study reveals the contribution of SCM to bottom hypoxia off the PRE, which is unreported and likely occurs in other coastal hypoxic systems.

Development of subsurface chlorophyll maximum layer and its contribution to the primary productivity of water column in a large subtropical reservoir

The phenomenon of subsurface chlorophyll maximum (SCM) layer emerging at a certain water depth is commonly found in stratified water bodies. Also, it is a crucial contributing region to the primary productivity of the water column. Currently, there is a lack of concern about the occurrence of SCM phenomena in studies targeting inland water bodies such as natural lakes and artificial reservoirs. This led to a significant underestimation of the level of primary productivity in these water bodies and their trophic state. In this study, a subtropical reservoir (the Xinanjiang Reservoir, XAJR) was investigated, to understand the characteristics of SCM layer in deep-large reservoir and its contribution to the primary productivity of the water column. Water sampling were conducted from September 2020 to August 2021, and in September 2022. Buoy station data for this reservoir between 2019 and 2021 were also collected. Based on the detailed observations of the water column profile in riverine area (X1), transitional area (X2), and central area (X3 and X4) of this reservoir, it was found that there was an obvious SCM phenomenon, which was closely related to the characteristics of seasonal thermal stratification. The SCM layer of XAJR appeared at depth around 3–5 m underwater from May to August, and as the thermal stratification strength increased, so did the depth and thickness of the SCM layer. It was estimated that gross primary productivity of euphotic layer of XAJR ranged from 347.9 to 4508.6 mgC·m−2·d−1. The average primary productivity level of the SCM layer reached 1411.7mgC·m−2·d−1, accounting for about 40–90% of the gross primary productivity of euphotic layer. This study contributes to a better understanding of the factors influencing changes in the development of the SCM layer in large reservoirs, as well as its critical role in the inland water carbon cycle.

The emergence and dominance of Planktothrix rubescens as an hypolimnetic cyanobacterium in response to re‐oligotrophication of a deep peri‐alpine lake

AbstractToxic cyanobacteria, such as Planktothrix rubescens, came to dominate temperate lakes in the mid‐to‐late 20th century, as a result of eutrophication. Even after decades of re‐oligotrophication, where phosphorus levels were reduced by 1–2 orders of magnitude, P. rubescens remains present in various lakes. In this study, we examine the persistence and changes of P. rubescens deep chlorophyll maximum (DCM) in Lake Hallwil (Switzerland) over 35 yr of steadily decreasing phosphorous concentrations. Although lake transparency increased and the euphotic layer deepened during this period, the P. rubescens population maximum moved even deeper. It is now found ca. 7.7 m deeper than its shallowest position in the 2000s, and this depth no longer coincides with the depth of maximal water column stability. P. rubescens neutral buoyancy has now driven it beneath the stable metalimnion into the hypolimnion, where buoyancy regulation is restricted due to reduced metabolic activity at low light and low temperature. If P. rubescens DCM continues to deepen each year, it will eventually reach a region of lower stability in the hypolimnion where turbulent conditions are strong enough to disperse the DCM. We also explore the mechanisms that ensure P. rubescens ongoing presence in peri‐alpine lakes despite strong re‐oligotrophication and ongoing climate change. We find that P. rubescens in the lake is mainly sustained by growth during fully mixed conditions in winter, not during stratification in summer. This may contradict another commonly made prediction that periods of longer stratification will promote future blooms of this cyanobacterium.

Frontogenesis Elevates the Maximum Chlorophyll a Concentration at the Subsurface Near the Kuroshio During Well‐Stratified Seasons

AbstractThe contribution that fronts and frontogenesis make to subsurface chlorophyll a (CHL) concentration was investigated near the Kuroshio using data from multiple surveys across the Kuroshio during summer and fall and satellite sea surface height (SSH) data alongside high‐resolution reanalysis data set. An intensive survey was conducted during a period of frontogenesis as seen from the in‐situ and satellite‐derived velocity data as well as the analysis of the reanalysis data set. Physical and biological observation showed shoaling of the nitracline and a subsurface CHL maximum (SCM) near the evolving front. The CHL concentration at the SCM (CHLSCM) was large near the front compared with the concentration in ambient water. Sections of 10 repeat seasonal surveys also captured the frontal structure, the shoaling of the nitracline, and the increase in CHLSCM as well as their interrelationship. The CHLSCM was large, and the depth of the SCM (ZSCM) was shallow above a large horizontal density gradient (i.e., marked mesoscale frontal structure) in the ocean interior. In addition, we demonstrate that the lateral strain rate, which is a measure of frontogenesis obtained from the SSH, can be used to estimate ZSCM and CHLSCM. Frontogenesis, secondary circulation, symmetric instability, and vertical diffusion were considered to cause the nutrient supply to the subsurface layer. The surveys and analysis indicate that fronts and frontogenesis increase CHLSCM, and the physical data obtained from both in‐situ and satellite observations can be used to estimate CHL distribution in the ocean interior.

Rhodopsin-mediated nutrient uptake by cultivated photoheterotrophic Verrucomicrobiota

Rhodopsin photosystems convert light energy into electrochemical gradients used by the cell to produce ATP, or for other energy-demanding processes. While these photosystems are widespread in the ocean and have been identified in diverse microbial taxonomic groups, their physiological role in vivo has only been studied in few marine bacterial strains. Recent metagenomic studies revealed the presence of rhodopsin genes in the understudied Verrucomicrobiota phylum, yet their distribution within different Verrucomicrobiota lineages, their diversity, and function remain unknown. In this study, we show that more than 7% of Verrucomicrobiota genomes (n = 2916) harbor rhodopsins of different types. Furthermore, we describe the first two cultivated rhodopsin-containing strains, one harboring a proteorhodopsin gene and the other a xanthorhodopsin gene, allowing us to characterize their physiology under laboratory-controlled conditions. The strains were isolated in a previous study from the Eastern Mediterranean Sea and read mapping of 16S rRNA gene amplicons showed the highest abundances of these strains at the deep chlorophyll maximum (source of their inoculum) in winter and spring, with a substantial decrease in summer. Genomic analysis of the isolates suggests that motility and degradation of organic material, both energy demanding functions, may be supported by rhodopsin phototrophy in Verrucomicrobiota. Under culture conditions, we show that rhodopsin phototrophy occurs under carbon starvation, with light-mediated energy generation supporting sugar transport into the cells. Overall, this study suggests that photoheterotrophic Verrucomicrobiota may occupy an ecological niche where energy harvested from light enables bacterial motility toward organic matter and supports nutrient uptake.

Response of coccolithophore communities to oceanographic and atmospheric processes across the North- and Equatorial Atlantic

Changes in coccolithophore productivity in response to climate-driven ocean warming are likely to have cascading biogeochemical effects that feed back to the changing climate. This paper investigates the role (and interplay) of large-scale oceanographic and atmospheric processes across the North- and Equatorial Atlantic, including Saharan dust deposition, on the distribution of coccolithophore communities. The study is based on biological and hydrological data collected across the photic zone of the ocean, and aerosol data collected from the lower atmosphere, across 50°N–1°S during the Atlantic Meridional Transect in boreal Autumn of 2018 (AMT28), in synergy with Earth Observations. Results confirm existing understanding of the distribution of coccolithophore communities which are related to major meridional hydrological gradients across the North Atlantic. Dynamic, oxygenated and microphytoplankton-enriched waters at higher-latitudes were characterized by less diverse coccolithophore populations, dominated by placolith-bearing r-selected coccolithophores. In contrast, the heavily stratified and picoplankton-enriched waters of the subtropical gyre revealed more diverse populations, dominated by umbelliform coccolithophores and holococcolithophores at the surface, and by floriform taxa in the lower photic zone. Mean concentrations of 14.4×103cells/L present in the North Atlantic Tropical Gyre Province (30–12°N), only slightly lower compared to 17.7×103cells/L produced in the North Atlantic Drift province (50–40°N), provide a snapshot perspective on the importance of coccolithophore production in heavily stratified gyre conditions. Higher concentrations of 19’-Hexanoyloxyfucoxanthin (HexFuco) in regions of enhanced production of r-selected placolith-bearing species suggest that this pigment should not be generalized as a proxy for the entire coccolithophore community. Enhanced abundances of fast-bloomingEmiliania huxleyiandGephyrocapsa oceanica, and of cyanobacteria (including both picoplankton and N2-fixingTrichodesmiumspp.) at the surface of the region of more persistent Saharan dust deposition (at ~12-10°N) appeared to result from dust-born nutrient input. Underneath this stratified surface layer, enhanced productivity in the deep chlorophyll maximum (DCM) appeared decoupled from that on the surface, fueled by geostrophic eastward shoaling of the nutricline across the tropical North Atlantic. As this was the region of highest macronutrient concentrations measured along and below the nutricline, our data suggest that the NE tropical Atlantic may act as a permanent dust-born nutrient depocenter as previously hypothesized.